Pseudotomentella umbrina (Fr.) M.J.Larsen, Canad. J. Bot. 45: 1298 (1967)

Svantesson, Sten, Larsson, Karl-Henrik, Koljalg, Urmas, W. May, Tom, Patrik Cangren,, Henrik Nilsson, R. & Larsson, Ellen, 2019, Solving the taxonomic identity of Pseudotomentellatristis s. l. (Thelephorales, Basidiomycota) - a multi-gene phylogeny and taxonomic review, integrating ecological and geographical data, MycoKeys 50, pp. 1-77 : 43-48

Pseudotomentella umbrina (Fr.) M.J.Larsen, Canad. J. Bot. 45: 1298 (1967) Fig. 18

Homotypic names.

Thelephora umbrina Fr. Elench. fung. 1: 199 (1828), non Pers. (1801), sanctioned name [Fries explicitly excluded T. umbrina Pers. from his concept]. Hypochnus umbrinus (Fr.) Fr. [basionym not cited], Summa veg. Scand.: 337 (1849), non Wallr. (1833), illegitimate name [combination also made by Quélet (1888) and Burt (1916)]. Corticium umbrinum (Fr.) Fr., Hymenomyc. eur.: 658 (1874). Coniophora umbrina (Fr.) Sacc., Syll. fung. 6: 652 (1888) [as (Alb. & Schwein.) Fr.]. Tomentella umbrina (Fr.) Litsch., Bull. Soc. Mycol. France. 49(1): 52 (June 20, 1933) [combination also made by Donk, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht. 9: 29 (before July 7 1933)]. Prillieuxia umbrina (Fr.) Park.-Rhodes 1956 Ann. Bot. (Oxford). 20(78): 258. 1956, invalid name, basionym not cited. Tomentellastrum umbrinum (Fr.) Svrček, Ceská Mykol. 12(2): 70 (1958).

Type.

SWEDEN. Småland: Femsjö, E. Fries (neotype: Herb. Fries [UPS F003106]!, designated by E.A. Burt in Ann. Missouri Bot. Gard 3: 213 (1916)); Småland: Hylte, Femsjö, Femsjö Church Nature Reserve, boreonemoral, mixed forest on soil with intermediate pH, 7 September 2016, S. Svantesson 351 (EPITYPE: GB!, here designated, MycoBank Typification No. MBT384818, GenBank Acc. No. ITS: MK290700).

UNITE SH.

SH030549.07FU

Description.

Basidiomata annual, resupinate, membranaceous, effused - often to several tens of centimetres in diameter. Mature parts continuous, with a cottony texture when fresh and a rather firm, fibrous and compact, yet quite soft and elastic texture when dried. Hymenium smooth, but sometimes strongly undulating; blue grey or purplish-grey to pale brown or brown when fresh, pale brown to brown when dried, sometimes with a reddish or greyish hue. Immature parts discontinuous, byssoid with a cottony texture, both when fresh and when dried. Subhymenium and hymenium of immature parts pale blue grey or pale purplish-grey to pale brown when fresh, pale brown when dried. Subiculum well developed, loose, fibrous, orange brown; often forms the outer edge of basidiomata, extending noticeably beyond the hymenium.

Hyphal cords lacking, but loose bundles of subicular hyphae sometimes present.

Hyphal system monomitic, clamp connections and reaction in Melzer’s reagent absent from all hyphae.

Subicular hyphae noticeably long and straight, thick-walled; forming a loose tissue. Individual hyphae 3.3-4.8 (-5.3) μm wide, with a mean width of 4.0-4.3 μm; orange brown to brown in KOH, orange brown in water.

Subhymenial hyphae often somewhat sinuous, thin to thick-walled; forming a rather dense tissue. Individual hyphae (2.8-) 2.9-5.0 (-5.9) μm wide, with a mean width of 3.7-4.2 μm; in the upper parts, pale green in KOH, sometimes with a faintly blue or brown hue; in the lower parts, orange brown to brown; in water, orange brown, with strongly granular contents.

Encrustation lacking.

Basidia with four slightly curved sterigmata, occasionally two-sterigmate; clavate to narrowly clavate, sometimes clavopedunculate, thin-walled, with one-three slight constrictions. Dimensions: (54-) 57-71 (-77) × (8.3-) 8.5-10.9 (-12.4) μm; mean dimensions: 60-64 × 9.5-10.3 μm. Sterigmata (8.7-) 8.8-11.1 (-11.7) μm long, with a mean length of 9.6-10.5 μm. Colour for the great majority very pale green in KOH, sometimes with a faintly blue or brown hue (but not the blue green reaction present in other species), for a small number formed directly from subicular hyphae brown; sometimes with granular contents; in water orange brown and with strongly granular contents.

Cystidial organs lacking.

Basidiospores in frontal face generally with a broadly subellipsoid, triangular or subcircular basic shape and an unlobed, angular, nodulose or sometimes cross-shaped outline, covered in bi- or trifurcate, occasionally singularly attached, echinuli. A majority of the spores normally with three-six indistinct corners to distinct, square lobes; broadly ellipsoid, unlobed spores infrequently occurring (but dominate in some collections), as well as abnormally large spores originating from two-sterigmate basidia. Frontal dimensions: 7.7-9.3 (-9.4) × (7.6-) 7.9-9.1 (-9.4) μm; mean dimensions: 8.3-8.7 × 8.4-8.7 μm; Q-value: (0.9-) 1.0-1.1; mean Q-value: 1.0. Echinuli (0.7-) 0.8-1.5 μm long, with a mean length of 1.1-1.2 μm. Lateral face ellipsoid to narrowly ovoid or sometimes semicircular in shape, usually with evenly rounded edges, sometimes with one-three lobes. Lateral dimensions: 8.0-9.3 (-9.6) × (5.1-) 5.6-6.7 (-6.9) μm; mean dimensions: 8.4-8.7 × 6.0-6.1 μm; Q-value: (1.2-) 1.3-1.6 (-1.7); mean Q-value: 1.4-1.5. Colour in KOH pale green to pale brown; in water orange brown to brown; inamyloid.

Chlamydospores lacking.

Habitat.

P. umbrina has a wide ecological amplitude. Recent Scandinavian collections have been made in young to old deciduous, mixed and coniferous forests on soil with low to high pH, as well as on the tundra. The species has been found to form ectomycorrhiza with at least Abies alba , Alnus rubra , Betula nana , Betula pubescens ssp. czerepanovii, Betula pubescens ssp. pubescens, Dryas octopetala , Fagus sylvatica , Picea abies , Picea glauca , Picea mariana , Pinus banksiana , Pinus pinaster , Pinus sylvestris , Pseudotsuga menziesii , Pyrola media , Quercus petraea , Salix polaris and Tsuga canadensis (collection data; Kõljalg et al. 2005, Nilsson et al. 2019).

Distribution.

Basidiomata encountered in: Canada, Estonia, Finland, Norway, Sweden and the United Kingdom. Soil or root tip samples confirm presence also in: France, Poland, Spain and the United States.

Remarks.

The nomenclatural situation surrounding P. umbrina is complex. Fries described Thelephora umbrina , explicitly excluding Thelephora umbrina Pers., but synonymising it with Thelephora umbrina var. lignatilis Alb & Schwein. These names might represent different species or not, but in either case do not threaten Thelephora umbrina Fr., due to the sanctioning.

A large number of names synonymous with P. umbrina have been illegitimately or superfluously published. Fries himself (1847), as well as Quélet (1888) and Burt (1916), seem to have overlooked Wallroth’s (1833) combination of Hypochnus umbrinus (Alb. & Schwein.) Wallr. from Himantia umbrina Alb. & Schwein, (1805) and hence created illegitimate name combinations. The status of Donk’s (1933) combination of Tomentella umbrina (Fr.) Donk versus Litschauer’s (1933) remains hard to evaluate due to the fact that, although 20 June is known to be the date of Litschauer’s publication, 7 Julyis when Donk defended the thesis wherein he published his combination; the publication date of Donk’s thesis was probably at an unknown point in time prior to that of his dissertation. Combinations based on Thelephora umbrina where the combining authors cite Alb. & Schwein as authors of the basionym (e.g. Saccardo 1888) have to be considered miscitations, since there is no Thelephora umbrina Alb. & Schwein.

Our interpretation of Thelephora umbrina Fr. as the basionym follows Burt (1916), Litschauer (1933), Svrcek (1958) and Larsen (1967), although Rogers and Jackson (1943) considered it to be a synonym of Coniophora olivacea . The name is not used here in the sense of what we today interpret as C. olivacea , but in the sense of Burt’s (1916) type selection.

The material in the Fries Herbarium cited by Burt (1916), as the type of Thelephora umbrina Fr., constitutes a collection made by Fries at locus classicus, Femsjö, but in Fries’s own handwriting, it is identified as Corticium umbrinum (Fr.) Fr., a name he combined T. umbrina to in 1874 ( Fries 1874). Therefore, Burt’s typification cannot be considered a lectotype, but must be regarded as a neotype. We here designate an epitype from Femsjö, which matches the neotype morphologically.

Within the P. tristis group, basidiomata of P. umbrina can be recognised by their brown colour - blue or green colours are completely absent from immature parts and from the subhymenium of mature parts - their soft, rather elastic texture after drying and their microcharacters. Pseudotomentella umbrinascens is very similar but has slightly different microcharacters (see key). Hypochnus rhacodium (only known from the type) is also similar but has hard, brittle basidiomata after drying.

Additional specimens studied.

CANADA. Newfoundland: Crooked Knife, mixed forest with Betula , Alnus and Picea , 99 m a.s.l., 10 September 2008, U. Kõljalg (TU 108084*);

ESTONIA. Põlva: Vastse-Kuuste, older Pinus-Picea mixed forest between Kiidjärve and Taevaskoja, near Maarja village, 22 September 2005, U. Kõljalg (TU 100329, 100339, 100340); Saare: Muhu, Kesselaid, Karjalasma forest, Picea abies forest, 28 August 1998, Erast Parmasto (TAAM 174051); Põlva: Vastse-Kuuste, coniferous forest with Pinus and Picea along road between Kiidjärve and Taevaskoja, east of Ahja river, 18 August 2005, U. Kõljalg (TU 100194); Viljandi: Pääsmä laas, Sooma National Park, on a fallen Betula trunk over Halliste river, 7 September 2000, U. Kõljalg (TU 108538);

FINLAND. Kanta-Häme: Lammi, Kotinen Virgin Forest, 10 September 2001, U. Kõljalg (TU 108742, 108743, 108744); Etelä-Häme, Ruovesi, Siikaneva swamp islands, 14 September 1999, U. Kõljalg (TAAM 159809, 159810); Satakunta: Ilkaalinen, under Picea log and mosses, 29 August 2010, U. Kõljalg (TU 115017); Varsinais-Suomi: Parainen, Kuggö, 24 October 2009, P. Kunttu (TU115344*);

NORWAY. Oppland, Dovre, Hjerkinn, low alpine vegetation under Salix phyllicifolia , Salix lapponica and Betula nana , on soil with low pH, 14 September 2014, S. Svantesson 216, 221* (GB); Akershus: Asker, Skaugumåsen, boreonemoral, mixed forest on moderately alkaline, moderately nutrient-rich ground under, 23 September 2010, S. Svantesson (O F110268*); Troms: Kvænangen, Kvænangselva, boreal mixed forest on soil with low pH, 31 August 2013, B. Larsson and K.-H. Larsson (O F110269); Ibidem, boreal, deciduous forest on soil with intermediate pH, 31 August 2013, B. Larsson and K.-H. Larsson (O F110270, F110271); Oppland: Dombås, Hjerkinnholen, boreal, mixed forest on soil with low pH, 30 September 2013, K.-H. Larsson (O F110272, F110273, F110274, F110275, F110276, F110277); Sogn og Fjordane: Leikanger, Flætene-Vesterheim, boreonemoral, mixed forest on soil with low pH, 2 October 2012, K.-H. Larsson and S. Svantesson (O F110278, F110279, F110280); Sogn og Fjordane: Eid, Eitrefjellet, deciduous forest on soil with high pH, 25 September 2013, K.-H. Larsson (O F110281); Oppland, Dovre, Grimsdalen, Storberget, subalpine Betula pubescens ssp. czerepanovii forest on soil with low pH, 26 August 2010, K.-H. Larsson and S. Svantesson (O F110282, F110283, F110284, F110285, F110286); Aust-Agder: Tvedestrand, Eidbo, boreonemoral, mixed forest on soil with intermediate pH, 10 September 2010, S. Svantesson and N. Svensson (O F110307); Aust-Agder: Åmli, Gangsei W, boreonemoral, mixed forest on soil with low pH, 09 September 2010, S. Svantesson and N. Svensson (O F110308); Telemark: Drangedal, Asgjerdstigfjellet, boreonemoral, deciduous forest on on soil with intermediate pH, 28 September 2010, S. Svantesson and N. Svensson (O F110309, F 110310); Vest-Agder: Mandal, Uføra, nemoral, deciduous forest on soil with high pH, 26 September 2012, K.-H. Larsson and S. Svantesson (O F110287); Sogn og Fjordane: Leikanger, Kvinnafossen, boreonemoral, mixed forest on soil with high pH, 2 October 2012, K.-H. Larsson and S. Svantesson (O F110288); Nord-Tröndelag: Grong, Gartlandselva, boreal, coniferous forest on soil with low pH, 27 August 2012, K.-H. Larsson (O F110289, F110290, F110291, F110292, F110293, F110294); Nordland: Saltdal, Nystadneslia, boreal, mixed forest on soil with intermediate pH, 24 August 2012, K.-H. Larsson (O F110295, F110296*); Telemark: Tokke, Dalen, Huvestad, boreonemoral, mixed forest on soil with high pH, 28 September 2010, S. Svantesson and N. Svensson (O F110311); Akershus: Nannestad, Tomte farm, 3 September 2004, U. Kõljalg (TU 100005, 100007); Telemark: Sauherad, E of Nordagutu, W slope of Bjørndalsfjell along path to Svanastøl, 24 September 2003, K.-H. Larsson 12094 (TU); Buskerud: Nes, Alungruken, 25 September 1997, J. Stokland (TU 115209*), Rogaland: Forsand, Rössdalen, on Salix sp., 14 October 1998, K. Hjortstam 17918 (K.-H. Larsson private collection);

SWEDEN. Lycksele Lappmark: Storuman, Blaiken N, boreal, mixed, old-growth forest on fertile, moderately alkaline ground, 26 August 2015, S. Svantesson 137 (GB); Västerbotten: Umeå, Stora Tuvan, older, boreal, mixed forest on soil with low pH, 28 August 2015, S. Svantesson 174*, 175 (GB); Lule Lappmark: Gällivare, Ritsem, subalpine Betula pubescens ssp. czerepanovii forest on soil with low pH, 11 August 2016, S. Svantesson 234, 239*, 240 (GB); Lule Lappmark: Jokkmokk, Slappejaure NO, middle alpine vegetation on soil with high pH, 14 August 2016, S. Svantesson 255, 256 (GB); Lule Lappmark: Jokkmokk, Unna Duvgge, low alpine vegetation on soil with intermediate pH, 15 August 2016, S. Svantesson 277 (GB); Lule Lappmark: Jokkmokk, Ajajaure N, low alpine vegetation on soil with high pH, 16 August 2016, S. Svantesson 279, 280* (GB); Halland: Kungsbacka, Släp, Särö Västerskog, old growth Pinus and Quercus forest, under a Pinus log, 1 October 1999 U. Kõljalg (TAAM 159818); Ångermanland; Sollefteå, Sörgraninge mångfaldspark, Språngsjöberget, 9 September 2014, K.-H. Larsson 16608 (GB); Västergötland: Alingsås, Simmenäshalvön, Gräskärr, on Picea , 5 October 2008, B. and K. Hjortstam 20311, 20332 (K.-H. Larsson private collection); Ibidem, on wood of Quercus on the ground, 13 September 2004, K. Hjortstam 18795 (K.-H. Larsson private collection); Ibidem, on Picea bark, 17 October 2001, K. Hjortstam 18531 (K.-H. Larsson private collection); Västergötland: Vårgårda, Nårunga, Sandviksås, on branch of Quercus robur , 8 November 2000, Björn Nordén (TU 115240);); Öland: Böda, Fagerör, under log of Pinus sylvestris , 15 October 2016, E. Larsson 387-16 (GB); Öland: Böda, Bryum Sandvik, under log of Pinus sylvestris , 15 October 2016, E. Larsson 384B-16 (GB);

UNITED KINGDOM OF GREAT BRITAIN AND NORTHERN IRELAND. Scotland, Invernesshire: Glen Strathfarrar National Nature Reserve, ancient Pinus sylvestris forest with a few oak trees, 14 September 2005, U. Kõljalg (TU 100304); Scotland, Morayshire: Culbin Forest, planted Pinus sylvestris forest on sand dunes, 13 September 2005, U. Kõljalg (TU 100292).